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Genomic analysis of Pyrginae Burmeister, 1878 (Lepidoptera: Hesperiidae Latreille, 1809) with an emphasis on the tribes Achlyodini Burmeister, 1878 and Carcharodini Verity, 1940 reveals many inconsistencies between the resulting phylogeny and the current classification. These problems are corrected by proposing new taxa, changing the ranks of others, or synonymizing them, and transferring species between genera. As a result, five subtribes, one genus, 20 subgenera, and one species are proposed as new: Cyclosemiina Grishin, new subtribe (type genus Cyclosemia Mabille, 1878), Ilianina Grishin, new subtribe (type genus Iliana E. Bell, 1937), Nisoniadina Grishin, new subtribe (type genus Nisoniades Hübner, [1819]), Burcina Grishin, new subtribe (type genus Burca E. Bell and W. Comstock, 1948), and Pholisorina Grishin, new subtribe (type genus Pholisora Scudder, 1872), all in Carcharodini; Lirra Grishin, new genus (type species Leucochitonea limaea Hewitson, 1868) in Pythonidina Grishin, 2019; Trifa Grishin, new subgenus (type species Tagiades jacobus Plötz, 1884), Tuberna Grishin, new subgenus (type species Pythonides contubernalis Mabille, 1883), Ebona Grishin, new subgenus (type species Quadrus eboneus E. Bell, 1947), Noctis Grishin, new subgenus (type species Achlyodes accedens Mabille, 1895), and Cyrna Grishin, new subgenus (type species Achlyodes cyrna Mabille, 1895) of Quadrus Lindsey, 1925; Liddia Grishin, new subgenus (type species Helias pallida R. Felder, 1869), Minna Grishin, new subgenus (type species Achlyodes minna Evans, 1953), and Thilla Grishin, new subgenus (type species Eurypterus later Mabille, 1891) of Eantis Boisduval, 1836; Torgus Grishin, new subgenus (type species Ouleus gorgus E. Bell, 1937) of Iliana E. Bell, 1937; Fenops Grishin, new subgenus (type species Cabares enops Godman and Salvin, 1894) of Polyctor Evans, 1953; Bezus Grishin, new subgenus (type species Pellicia bessus Möschler, 1877) and Macarius Grishin, new subgenus (type species Pellicia macarius Herrich-Schäffer, 1870) of Nisoniades Hübner, [1819]; Quadralis Grishin, new subgenus (type species Pterygospidea extensa Mabille, 1891) of Gorgopas Godman and Salvin, 1894; Menuda Grishin, new subgenus (type species Nisoniades menuda Weeks, 1902) and Narycus Grishin, new subgenus (type species Pythonides narycus Mabille, 1889) of Perus Grishin, 2019; Bovaria Grishin, new subgenus (type species Achlyodes cyclops Mabille, 1876), Sebia Grishin, new subgenus (type species Nisoniades eusebius Plötz, 1884), and Stolla Grishin, new subgenus (type species Pholisora balsa E. Bell, 1937) of Bolla Mabille, 1903; Vulga Grishin, new subgenus (type species Achlyodes vulgata Möschler, 1879) and Capilla Grishin, new subgenus (type species Helias aurocapilla Staudinger, 1876, currently a junior subjective synonym of Hesperia musculus Burmeister, 1875) of Staphylus Godman and Salvin, 1896; and Quadrus (Zera) vivax Grishin, new species (type locality in Brazil: Rio de Janeiro). The following 10 are subgenera, not genera or synonyms: Ouleus Lindsey, 1925 and Zera Evans, 1953 of Quadrus Lindsey, 1925; Atarnes Godman and Salvin, 1897 and Eburuncus Grishin, 2012 of Milanion Godman and Salvin, 1895; Pachyneuria Mabille, 1888 and Austinus O. Mielke and Casagrande, 2016 of Sophista Plötz, 1879; Hemipteris Mabille, 1889 and Mictris Evans, 1955 of Pellicia Herrich-Schäffer, 1870; and Hesperopsis Dyar, 1905 and Scantilla Godman and Salvin, 1896 of Staphylus Godman and Salvin, 1896. The following 7 are species, not subspecies: Quadrus (Ebona) cristatus (Steinhauser, 1989) (not Quadrus (Ebona) negrus (Nicolay, 1980)), Quadrus (Quadrus) ophia (A. Butler, 1870) (not Quadrus (Quadrus) lugubris (R. Felder, 1869)), Quadrus (Zera) gellius (Mabille, 1903) and Quadrus (Zera) servius (Plötz, 1884) (not Quadrus (Zera) hyacinthinus (Mabille, 1877)), Mimia pazana Evans,1953 (not Mimia phidyle (Godman and Salvin, 1894)), Polyctor (Polyctor) dagua Evans, 1953 (not Polyctor (Polyctor) polyctor (Prittwitz, 1868)), and Staphylus (Vulga) satrap Evans, 1953 (not Staphylus (Vulga) saxos Evans, 1953); and these 8 are species, not synonyms: Quadrus (Zera) menedemus (Godman and Salvin, 1894) (not Quadrus (Zera) tetrastigma (Sepp, [1847])), Pellicia (Pellicia) bilinea Mabille, 1889 (not Pellicia (Pellicia) dimidiata Herrich-Schäffer, 1870), Pellicia (Hemipteris) nema Williams and Bell, 1939 (not Pellicia (Pellicia) theon Plötz, 1882), Bolla (Bovaria) sodalis Schaus, 1913 (not Bolla (Bolla) imbras (Godman and Salvin, 1896)), Bolla (Bovaria) aplica (E. Bell, 1937) (not Bolla (Sebia) eusebius (Plötz, 1884)), Bolla (Sebia) chilpancingo (E. Bell, 1937) (not Bolla (Bolla) subapicatus (Schaus, 1902)), and Bolla (Stolla) madrea (R. Williams and E. Bell, 1940) and Bolla (Stolla) hazelae (Hayward, 1940) (not Bolla (Stolla) zorilla (Plötz, 1886)). The following 2 are junior subjective synonyms: Achlyodes erisichthon Plötz, 1884 of Quadrus (Zera) servius (Plötz, 1884) (not a subspecies of Quadrus (Zera) tetrastigma (Sepp, [1847]) and Staphylus subapicatus Schaus, 1902 of Bolla (Bolla) imbras (Godman and Salvin, 1896). Furthermore, we propose the following additional new genus-species combination: Gindanes homer (Evans, 1953), Gindanes nides (O. Mielke and Casagrande, 2002), Gindanes maraca (O. Mielke and Casagrande, 1992), Gindanes jenmorrisae (Shuey and Ramírez. 2022), Gindanes tullia (Evans, 1953), Gindanes herennius (Geyer, [1838]), Gindanes proxenus (Godman and Salvin, 1895), Gindanes parallelus (Mabille, 1898), Gindanes braga (Evans, 1953), Gindanes hampa (Evans, 1953), Gindanes rosa (Steinhauser, 1989), Gindanes neivai (Hayward, 1940), Gindanes mundo (H. Freeman, 1979), Gindanes eminus (E. Bell, 1934), Quadrus (Trifa) francesius Freeman, 1969, Quadrus (Trifa) ineptus (Draudt, 1922), Quadrus (Trifa) jacobus (Plötz, 1884), Quadrus (Tuberna) lancea (Hewitson, 1868), Quadrus (Ebona) pescada (E. Bell, 1956), Lirra pteras (Godman and Salvin, 1895), and Lirra limaea (Hewitson, 1868) (not Pythonides Hübner, 1819); Quadrus (Cyrna) zora (Evans, 1953) (not Bolla Mabille, 1903); Eantis later (Mabille, 1891) and Eantis haber (Mabille, 1891) (not Aethilla Hewitson, 1868); Iliana (Torgus) gorgus (E. Bell, 1937) and Iliana (Torgus) taurus (Evans, 1953) (not Eantis Boisduval, 1836); Bolla (Stolla) evemerus (Godman and Salvin, 1896), Bolla (Stolla) chlora (Evans, 1953), Bolla (Stolla) astra (R. Williams and E. Bell, 1940), Bolla (Stolla) balsa (E. Bell, 1937), Bolla (Stolla) tridentis (Steinhauser, 1989), Bolla (Stolla) esmeraldus (L. Miller, 1966), Bolla (Stolla) chlorocephala (Latreille, [1824]), and Bolla (Stolla) incanus (E. Bell, 1932) (not Staphylus Godman and Salvin, 1896). Finally, lectotypes are designated for Achlyodes servius Plötz, 1884 (type locality in Brazil: Rio de Janeiro), Pellicia theon Plötz, 1882 (type locality in South America), and Nisoniades eusebius Plötz, 1884 (type locality in Central America). Unless stated otherwise, all subgenera, species, subspecies, and synonyms of mentioned genera and species are transferred with their parent taxa, and others remain as previously classified.
ZooBank registration. http://zoobank.org/B9AFA1A9-8664-4F31-B4D9-ACF7780C7CC6
A century and a half since the time of Hewitson, we are experiencing a renaissance in species discovery fueled by whole genome sequencing. A large-scale genomic analysis of Hesperiidae Latreille, 1809 (Lepidoptera), including primary type specimens, reveals a deluge of species new to science. One hundred of them (one in a new genus) are described here from the New World (type localities are given in parenthesis): Drephalys (Drephalys) diovalis Grishin, new species (Ecuador: Napo), Euriphellus panador Grishin, new species (Ecuador: Esmeraldas), Euriphellus panamicus Grishin, new species (Panama: Panama), Cecropterus (Thorybes) viridissimus Grishin, new species (Ecuador: Zamora-Chinchipe), Cecropterus (Murgaria) dariensis Grishin, new species (Panama: Darien), Urbanus (Urbanus) mericuti Grishin, new species (Ecuador: Napo), Telegonus (Telegonus) pastus Grishin, new species (Panama: Panama), Autochton (Autochton) dora Grishin, new species (Ecuador: Pastaza), Astraptes centralis Grishin, new species (Panama: Colón), Aguna claxonica Grishin, new species (Ecuador: Napo), Aguna esmeralda Grishin, new species (Ecuador: Esmeraldas), Aguna lata Grishin, new species (Guyana), Ridens angulinea Grishin, new species (Peru: Cuzco), Pythonides lera Grishin, new species (Peru: Cuzco), Pythonides latemarginatus Grishin, new species (Panama: Panama), Gindanes variegatus Grishin, new species (Brazil: Mato Grosso), Milanion (Milanion) virga Grishin, new species (Brazil: Rondônia), Milanion (Milanion) furvus Grishin, new species (Panama: Panama), Milanion (Milanion) laricus Grishin, new species (Ecuador: Napo), Charidia ronda Grishin, new species (Brazil: Rondônia), Pseudodrephalys tinas Grishin, new species (Peru: Loreto), Pseudodrephalys argus Grishin, new species (Suriname: Para), Achlyodes calvus Grishin, new species (Brazil: Santa Catarina), Spioniades artemis Grishin, new species (Panama: Panama), Spioniades artemidoides Grishin, new species (Brazil: Santa Catarina), Myrinia orieca Grishin, new species (Ecuador: Orellana), Myrinia aragua Grishin, new species (Venezuela: Aragua), Myrinia maculosa Grishin, new species (Guatemala), Myrinia manchada Grishin, new species (Guyana), Polyctor (Fenops) lamperus Grishin, new species (Panama: Darien), Nisoniades (Nisoniades) lutum Grishin, new species (Mexico: Guerrero. ), Bolla (Stolla) vena Grishin, new species (Venezuela: Aragua), Staphylus (Vulga) vula Grishin, new species (Mexico: Veracruz), Staphylus (Vulga) vulga Grishin, new species (Panama: Darien), Staphylus (Staphylus) rotundalus Grishin, new species (Ecuador: Napo), Staphylus (Staphylus) yucatanus Grishin, new species (Mexico: Quintana Roo/Yucatan), Heliopetes (Heliopetes) lana Grishin, new species (Guatemala), Canesia ella Grishin, new species (Venezuela: Barinas), Paches (Paches) loxeca Grishin, new species (Ecuador: Morona-Santiago), Clito congruens Grishin, new species (Panama: Colón), Cycloglypha corax Grishin, new species (Brazil: Rio de Janeiro), Festivia peruvia Grishin, new species (Peru: Huánuco), Decinea notata Grishin, new species (Ecuador: Napo), Pompeius fuscus Grishin, new species (Brazil: Minas Gerais), Vernia clara Grishin, new species (Panama: Chiriquí), Oligoria (Oligoria) obtena Grishin, new species (Ecuador: Napo), Thespieus mandal Grishin, new species (Brazil: Rio de Janeiro), Psoralis (Saniba) magnamacus Grishin, new species (Panama: Darien), Alychna ayonis Grishin, new species (Ecuador: Napo), Wahydra banios Grishin, new species (Ecuador: Tungurahua), Wahydra cuzcona Grishin, new species (Peru: Cuzco), Cynea (Cynea) aureofimbra Grishin, new species (Ecuador), Cynea (Nycea) quada Grishin, new species (Ecuador: Napo), Cynea (Quinta) achirae Grishin, new species (Mexico: Tamaulipas), Eutus amazonicus Grishin, new species (Peru: Madre de Dios), Eutus incus Grishin, new species (Peru: Cuzco), Eutus septemaculatus Grishin, new species (Brazil: Mato Grosso), Godmia viridicapita Grishin, new species (Ecuador: Napo), Rhomba pulla Grishin, new species (Peru: Cuzco), Niconiades victoria Grishin, new species (Mexico: Tamaulipas), Lancephallus purpurus Grishin, new genus and new species (Guyana), Mnasicles (Remella) ecua Grishin, new species (Ecuador: Pichincha), Amblyscirtes (Amblyscirtes) aeratus Grishin, new species (Mexico: Oaxaca), Amblyscirtes (Mastor) chrysoplea Grishin, new species (Mexico: Oaxaca), Amblyscirtes (Mastor) chrysomisa Grishin, new species (Mexico: Chiapas), Amblyscirtes (Flor) meridus Grishin, new species (Mexico: Veracruz), Rectava chiriquensis Grishin, new species (Panama: Chiriquí), Cobalopsis adictys Grishin, new species (Panama: Veraguas), Cymaenes melaporphyrus Grishin, new species (Mexico: San Luis Potosí), Lerema (Morys) ecuadorica Grishin, new species (Ecuador: Pichincha), Saturnus obscurior Grishin, new species (Panama: Darien), Cantha zoirodicta Grishin, new species (Peru: Madre de Dios), Cantha meiodicta Grishin, new species (Peru: Madre de Dios), Phlebodes duplex Grishin, new species (Guatemala: Cayuga), Lychnuchus (Enosis) valle Grishin, new species (Colombia: Valle), Eutychide ochoides Grishin, new species (Peru: Cuzco), Dion bora Grishin, new species (Panama: Darien), Dion occida Grishin, new species (Peru: Madre de Dios), Eprius (Eprius) veledinus Grishin, new species (Ecuador: Pichincha), Radiatus panamensis Grishin, new species (Panama: Panama), Pheraeus pulcher Grishin, new species (Peru: Madre de Dios), Callimormus rades Grishin, new species (Panama: Panama), Gubrus lubens Grishin, new species (Ecuador: Loja), Ludens labens Grishin, new species (Panama: Darien), Rigga isa Grishin, new species (Ecuador: Napo), Flaccilla lactea Grishin, new species (Peru: Cuzco), Falga athena Grishin, new species (Panama: Darien), Panoquina jay Grishin, new species (Peru: Loreto), Calpodes salianus Grishin, new species (Peru: Madre de Dios), Calpodes stingo Grishin, new species (Ecuador: Sucumbíos), Aides nobra Grishin, new species (Panama: Colón), Thracides pavo Grishin, new species (Mexico: Tabasco), Talides eluta Grishin, new species (Peru: Cuzco), Talides laeta Grishin, new species (Peru: Cuzco), Neoxeniades angustior Grishin, new species (Brazil: Rio de Janeiro), Damas zea Grishin, new species (Guyana), Tromba xantha Grishin, new species (Mexico: Veracruz), Perichares fura Grishin, new species (Ecuador: Pichincha), Carystoides (Balma) goliath Grishin, new species (Colombia: Valle), and Agathymus galeana Grishin, new species (Mexico: Nuevo Leon). Additionally, we present evidence to support 22 taxa as species (not subspecies or synonyms) and synonymize one genus and four species. Namely, the following taxa are species: Milanion pilta Evans, 1953 (not Milanion pilumnus Mabille and Boullet, 1917), Milanion latior Mabille and Boullet, 1917 (not a synonym of Milanion marciana Godman and Salvin, 1895), Charidia pilea Evans, 1953, and Charidia pocus Evans, 1953 (not Charidia lucaria (Hewitson, 1868)), Paches (Paches) gloriosus Röber, 1925 and Paches (Paches) loxana Evans, 1953 (not Paches (Paches) loxus (Westwood, 1852)), Spioniades anta Evans, 1953 (not Spioniades abbreviata (Mabille, 1888)), Decinea onasima (Hewitson, 1877) and Decinea formosus (Hayward, 1940) (not Decinea dama (Herrich-Schäffer, 1869)), Thespieus guerreronis (Dyar, 1913) (not Thespieus dalman (Latreille, [1824])), Cynea (Nycea) erebina (Möschler, 1879) and Cynea (Nycea) cleochares (Mabille, 1891) (not Cynea (Cynea) diluta (Herrich-Schäffer, 1869)), Amblyscirtes (Mastor) repta Evans, 1955 (not Amblyscirtes (Flor) florus (Godman, 1900)), Saturnus tiberius (Möschler, 1883), Saturnus conspicuus (E. Bell, 1941), Saturnus meton (Mabille, 1891), and Saturnus obscurus (E. Bell, 1941) (not Saturnus reticulata (Plötz, 1883)), Phlebodes sifax Evans, 1955 (not Phlebodes campo (E. Bell, 1947)), Eutychide ochus Godman, 1900 and Eutychide rogersi (Kaye, 1914) (not a subspecies and a synonym, respectively, of Eutychide subcordata (Herrich-Schäffer, 1869)), Falga mirabilis Evans, 1955, Falga jacta Evans, 1955, and Falga ombra Evans, 1955 (not Falga jeconia (A. Butler, 1870)); and the following taxa are junior subjective synonyms: Libra Evans, 1955 (of Phemiades Hübner, [1819]), Papilio clito Fabricius, 1787 of Milanion hemes hemes (Cramer, 1777), Pamphila hycsos Mabille, 1891 of Cynea (Nycea) erebina (Möschler, 1879), Hesperia olympia Plötz, 1882 of Eutychide subcordata (Herrich-Schäffer, 1869), and Hesperia ocrinus Plötz, 1882 of Aides aegita (Hewitson, 1866). Furthermore, we propose new combinations for genus-species: Lychnuchus (Enosis) ponka (Evans, 1955) (not Thoon Godman, 1900), and species-subspecies: Charidia pocus mayo Evans, 1953 (not Charidia lucaria (Hewitson, 1868)), Decinea onasima boliviensis (E. Bell, 1930) (not Decinea dama (Herrich-Schäffer, 1869)), Cynea (Nycea) erebina somba Evans, 1955 (not Pamphila hycsos Mabille, 1891), Saturnus tiberius suffuscus (Hayward, 1940) (not Saturnus reticulata (Plötz, 1883)), and Falga mirabilis odol Evans, 1955 (not Falga jeconia (A. Butler, 1870)). Then, Milanion pilumnus var. hemestinus Mabille and Boullet, 1917 is a junior subjective synonym of Milanion pilumnus pilumnus Mabille and Boullet, 1917, not of Milanion leucaspis (Mabille, 1878). Lectotypes are designated for nine taxa (names in original combinations below): Pellicia bromias Godman and Salvin, 1894 (Mexico: Veracruz, Atoyac), Nisoniades perforata Möschler, 1879 (Colombia), Helias ascalaphus Staudinger, 1876 (central Panama), Pamphila hycsos Mabille, 1891 (Colombia), Amblyscirtes fluonia Godman, 1900 (Mexico: Guerrero, Xocomanatlan), Mastor anubis Godman, 1900 (Mexico: Guerrero, Omiltemi), Eutychide ochus Godman, 1900 (Mexico: Veracruz, Atoyac), Cobalus subcordata Herrich-Schäffer, 1869 (Southeast Brazil), and Thracides xanthura Godman, 1901 (Panama: Chiriquí Province, Bugaba). A neotype is designated for Eudamus briccius Plötz, 1881 (Guyana: Iwokrama Forest).
ZooBank registration. urn:lsid:zoobank.org:pub:ACDF923B-906D-460E-9707-259E0ECDBCA8